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Zinc is an essential trace metal for oceanic primary producers with the highest concentrations in polar oceans. However, its role in the biological functioning and adaptive evolution of polar phytoplankton remains enigmatic. Here, we have applied a combination of evolutionary genomics, quantitative proteomics, co-expression analyses and cellular physiology to suggest that model polar phytoplankton species have a higher demand for zinc because of elevated cellular levels of zinc-binding proteins. We propose that adaptive expansion of regulatory zinc-finger protein families, co-expanded and co-expressed zinc-binding proteins families involved in photosynthesis and growth in these microalgal species and their natural communities were identified to be responsible for the higher zinc demand. The expression of their encoding genes in eukaryotic phytoplankton metatranscriptomes from pole-to-pole was identified to correlate not only with dissolved zinc concentrations in the upper ocean but also with temperature, suggesting that environmental conditions of polar oceans are responsible for an increased demand of zinc. These results suggest that zinc plays an important role in supporting photosynthetic growth in eukaryotic polar phytoplankton and that this has been critical for algal colonization of low-temperature polar oceans.

Since the article “Primary Production, an Index of Climate Change in the Ocean: Satellite-Based Estimates over Two Decades” by Kulk et al. [1] was published, we discovered an error in the code of the primary production model, which crept in when the code was updated from the original version described by Platt and Sathyendranath (1988), Sathyendranath et al. (1995) and Longhurst et al. (1995) ([2,31,52] in [1]). The main error in the code led to a time interval for the integration of daily water-column primary production that was shorter than it should have been. As a consequence, daily surface irradiance and hence primary production were systematically underestimated by 20–25% for the entire time series. We also discovered that the Photosynthetic Active Radiation (PAR) products of the National Aeronautics and Space Administration (NASA) that were used to scale the daily light cycle were rounded down for 2003–2019 (MODIS years), which led to an additional but small underestimation of daily surface irradiance. In addition to addressing these errors, we have included a merged time series of the PAR product to remove inter-sensor biases (as described in the corrected text of Appendix B; see below). The main corrections increased our estimate of global annual primary production on average by +23.9% between 1998 and 2018, while the correction of the rounding error in the PAR products increased global annual primary production between 2003 and 2018 by +0.9%. Inclusion of the merged PAR product in the primary production model caused a −0.25% decrease in global annual primary production between 1998 and 2002 and a +0.08% increase between 2003 and 2010 (relative to the aforementioned +23.9% increase for the entire time series). Our estimate of global annual primary production between 1998 and 2018 now is 48.7 to 52.5 Gt C y−1 instead of the published estimate of 38.8 to 42.1 Gt C y−1. Although this is a substantial increase in the estimate of primary production, the results

Primary production by marine phytoplankton is one of the largest fluxes of carbon on our planet. In the past few decades, considerable progress has been made in estimating global primary production at high spatial and temporal scales by combining in situ measurements of primary production with remote-sensing observations of phytoplankton biomass. One of the major challenges in this approach lies in the assignment of the appropriate model parameters that define the photosynthetic response of phytoplankton to the light field. In the present study, a global database of in situ measurements of photosynthesis versus irradiance (P-I) parameters and a 20-year record of climate quality satellite observations were used to assess global primary production and its variability with seasons and locations as well as between years. In addition, the sensitivity of the computed primary production to potential changes in the photosynthetic response of phytoplankton cells under changing environmental conditions was investigated. Global annual primary production varied from 38.8 to 42.1 Gt C yr −1 over the period of 1998–2018. Inter-annual changes in global primary production did not follow a linear trend, and regional differences in the magnitude and direction of change in primary production were observed. Trends in primary production followed directly from changes in chlorophyll-a and were related to changes in the physico-chemical conditions of the water column due to inter-annual and multidecadal climate oscillations. Moreover, the sensitivity analysis in which P-I parameters were adjusted by ±1 standard deviation showed the importance of accurately assigning photosynthetic parameters in global and regional calculations of primary production. The assimilation number of the P-I curve showed strong relationships with environmental variables such as temperature and had a practically one-to-one relationship with the magnitude of change in primary production. In the future, such empiric

Cabled coastal observatories are often seen as future-oriented marine technology that enables science to conduct observational and experimental studies under water year-round, independent of physical accessibility to the target area. Additionally, the availability of (unrestricted) electricity and an Internet connection under water allows the operation of complex experimental setups and sensor systems for longer periods of time, thus creating a kind of laboratory beneath the water. After successful operation for several decades in the terrestrial and atmospheric research field, remote controlled observatory technology finally also enables marine scientists to take advantage of the rapidly developing communication technology. The continuous operation of two cabled observatories in the southern North Sea and off the Svalbard coast since 2012 shows that even highly complex sensor systems, such as stereo-optical cameras, video plankton recorders or systems for measuring the marine carbonate system, can be successfully operated remotely year-round facilitating continuous scientific access to areas that are difficult to reach, such as the polar seas or the North Sea. Experience also shows, however, that the challenges of operating a cabled coastal observatory go far beyond the provision of electricity and network connection under water. In this manuscript, the essential developmental stages of the “COSYNA Shallow Water Underwater Node” system are presented, and the difficulties and solutions that have arisen in the course of operation since 2012 are addressed with regard to technical, organizational and scientific aspects.

Effects of prolonged darkness on Arctic phytoplankton composition were investigated with lab experiments and a pigment time series in Kongsfjorden, Spitsbergen (78°55′N). Chlorophyll a (Chl a), pigment composition, particulate organic carbon, cell numbers, and photosynthetic characteristics were studied in Arctic diatoms (Thalassiosira antarctica, Thalassiosira nordenskioeldii) and flagellates (Rhodomonas sp., Micromonas sp.) during 8 weeks of darkness and subsequent recovery in irradiance. Loss of photosynthetic functionality after 2 weeks of darkness was reversible in all species when returned to irradiance. Diatoms were more resistant to prolonged darkness (> 2 weeks) compared to the flagellates, with lower decline rates of Chl a and maximum quantum yield of PSII. T. nordenskioeldii showed rapid growth during recovery throughout 8 weeks of dark incubation, whereas recovery of flagellates diminished within 4 weeks. Ratios of taxonomic marker pigments relative to Chl a of all species showed limited variation during 8 weeks of dark incubation. The experimentally observed enhanced dark survival of diatoms was in agreement with pigment observations during four polar nights (2013–2017) in Kongsfjorden, which showed increased relative diatom abundance during declining biomass (down to 0.02 mg Chl a m−3). Therefore, a period of prolonged darkness gives Arctic diatoms a head start during the early stages of the spring bloom. The taxon‐specific survival traits can influence the geographical distribution of diatoms and flagellates within the polar oceans and their phenology. Furthermore, the persistence of Chl a of nonviable phytoplankton during darkness might influence biomass estimates during the polar night.

Phytoplankton biomass and composition was investigated in a high Arctic fjord (Kongsfjorden, 79◦N, 11◦40′E) using year round weekly pigment samples collected from October 2013 to December 2014. In addition, phytoplankton dynamics supplemented with physical and chemical characteristics of the 2014 spring bloom (April–June 2014) were assessed in two locations in Kongsfjorden. The goal was to elucidate effects of Atlantic advection on spatial phytoplankton chlorophyll-a (chl-a) and taxonomic composition. Chl-a declined during the polar night to a minimum of 0.01mg m−3, followed by a 1000-fold increase until May 28. Atlantic advection prevented sea ice formation and increased springtime melting of marine terminating glaciers. This coincided with spatial and temporal differences in abundances of flagellates (prasinophytes, haptophytes, cryptophytes, and chrysophytes) and diatoms in early spring. More flagellated phytoplankton were observed in the non-stratified central Kongsfjorden, whereas diatoms were more abundant in the stratified inner fjord. Contrasting conditions between locations were reduced when glacial melt water stratification expanded toward the mouth of the fjord, mediating a diatom dominated surface bloom at both locations. We suggest that glacial melt water governs spring bloom spatial timing and composition in the absence of sea ice driven stratification. The spring bloom exhausted surface nutrient concentrations by the end of May. The nutrient limited post bloom period (June–October) was characterized by reduced biomass and pigments of flagellated phytoplankton, consisting of prasinophytes, haptophytes, chrysophytes, and to a lesser extent cryptophytes and peridinin-containing dinoflagellates.